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Evidently it is still widely held
that pterosaurs appeared suddenly without antecedent. As evidence of this failure to follow the data, I came across a blog called Tetrapod Flight in which the author, Leon Linde, writes on Monday, March 16, 2015:

The first tetrapods to evolve powered flight were the pterosaurs. True

These were a group of archosaurs related to the dinosaurs, but not dinosaurs themselves. False. Pterosaurs are lepidosaurs, not related to dinos.

The earliest known pterosaur was Eudimorphodon, who lived in what is now Italy around 230-220 million years ago, in the late Triassic. True enough

However, while the earliest known pterosaur, Eudimorphodon had specialised multi-cusped teeth not found in any of the later pterosaurs, so it would not have been ancestral to them but rather part of a distinct pterosaur lineage that died out in the Triassic. False. Multicupsed teeth are found in several Triassic pterosaurs AND in their proximal sisters.

Furthermore, both Eudimorphodon and other late Triassic pterosaurs are “completely” developed, having all the typical pterosaur skeletal characteristics. True. That’s why they are called pterosaurs. They have all ‘the goods’.

This suggests the origins of pterosaurs may lie even further back in the past, in the earlier Triassic or perhaps even in the Permian (Wellnhofer, 1991). Yes to the earlier (Middle) Triassic (Cosesaurus) and No to the Permian.

No fossils of the pterosaurs’ immediate ancestors are known. False. We have pterosaur proximal and distant ancestors going back to basal tetrapods with fins. Click here.

The most likely theory on their origins is that they evolved from arboreal creatures that would leap from branch to branch, flapping their forelimbs to stay airborne longer. Actually we have evidence for this scenario chronicled here.

Pterosaur hips had great freedom of movement, their knees and ankles were hinge-like and their feet were plantigrade. True, True, True and False. Some beachcombers had plantigrade feet, but basal forms did not.

The knees and ankles did not permit the necessary rotation for them to move bipedally, so pterosaurs were obligate quadrupeds (though they may have had bipedal ancestors). False. Like living bipedal lizards, basal pterosaurs were bipedal and agile. We have their tracks! Later forms, especially beachcombers, were quadrupedal, and we have their tracks, too.

A possible explanation for these features is that the early pterosaurs or proto-pterosaurs were arboreal creatures that evolved powerful leaping from branch to branch as an active mode of transport not dissimilar to that of arboreal leaping primates (Christopher, 1997). This reference should be Bennett 1997. Powerful leaping, fast running, yes, but without the use of the hands, which were flapping like those of birds and getting larger. Hard to develop wings when you’re using your hands on the ground.

These arboreal leapers would not have been gliders, who merely fall slowly downwards and forwards with the help of special flaps, but rather creatures utilising a quite different form of locomotion, one that led them to eventually having their forelimbs evolve into more and more sophisticated flapping airfoils. True! But not like the image below (which appeared on the blog post). This pretty but bogus image shows no flapping and no reason or benefit to having proto-wings on this dinosaur that should have been a fenestrasaur.

Sadlythis state of affairs in pterosaur research shows that the general public and pterosaur artists and workers alike are still stuck in the tail-dragging age. Evidently, they have decided to shun and overlook recent data that chronicles and documents the origin of pterosaurs. See below and here.

Arcticodactylus cromptonellus (Kellner 2015, originally Eudimorphodon cromptonellus Jenkins et al. 1999, 1999; MGUH VP 3393) Late Triassic ~210mya ~8 cm snout to vent length was a tiny pterosaur derived from a sister to Eudimorphodon ranzii and phylogenetically preceded Campylognathoides and BSp 1994 specimen attributed to Eudimorphodon. Whether it was a juvenile or a tiny adult cannot be determined because juveniles and even embryos are identical to adults in pterosaurs. Note that that rostrum was not shorter and the orbit was not larger than in sister taxa. This specimen is one of the smallest known pterosaurs., but not THE smallest (Fig. 1) contra the Wikipedia article. That honor goes to B St 1967 I 276.

Figure 1. Articodactylus is evidently NOT the smallest pterosaur. That honor still goes to an unnamed specimen (not a Pterodactylus kochi juvenile) B St 1967 I 276.

Distinct from E. ranzii,
the skull of Arctiodactylus had a rounder, less triangular orbit. The jugal was not as deep. The sternal complex did not have small lateral processes. The humerus was not as robust. The fingers were longer an more gracile. The prepubis was distinctly shaped.

Distinct fromBergamodactylus the femur and tibia were smaller but the metatarsals were longer, compact and nearly subequal in length with IV smaller than III.

No current discoveries are found in the latest textbooks.
That’s because it takes time (years typically) for textbooks to be (in reverse order) assigned, accepted, distributed, printed, edited, written and illustrated, researched and concepted. Textbook publishers are out to sell the maximum number of books, so they write to the current consensus, which may be in flux on several points and hypotheses. The current consensus may also be wrong–but it remains the consensus.

There are no courses
at any colleges entitled, PTEROSAURS 101, 102 or 103. Who would attend? There are only two dozen people in the world who have an interest, who study them, or contribute to what we know about them. And where is the consensus? On some points, there is no consensus!! And all too often “the consensus” is holding on to outmoded, invalid and unverifiable paradigms (see below).

Every new fossil specimen is really a new chapter
in an ever expanding textbook on paleontology. And all paleontologists who publish are contributing authors to that future textbook.

Striiving for veracity
It is important for all workers to see things as they are in specimens, and not to reinterpret them to fit an established paradigm, no matter the temptation to do otherwise. For instance, narrow chord wing preservation in pterosaurs is not the result of ‘shrinkage’ as some workers report. Rather it is what it is, universal. All pterosaur specimens have narrow chord wings. If you know one that is different, please tell me. I know one that appears different, but that’s because part of its arm was ripped away and displaced. Look closely. That’s the way it is.

If Galileo
went to school as a teenager and found the following question on a test: “If object A at ten pounds and object B at 10 ounces both fall from 1000 feet at the precisely the same moment, how many seconds ahead of B will A strike the ground?” He’d would not have even had the opportunity to choose answer E. “zero seconds.” Common knowledge at the time, based on Aristotle, would not have allowed it, no matter the facts of this case, proven by experiment. This went on for centuries.

Similarly,
if you were in college today and were given the multiple choice question, “Which one of the following taxa is most closely related to pterosaurs? A. Dinosaurs. B. Scleromochlus. C. Proterochampsids (including Lagerpeton). D. Euparkeria. E. Erythrosucids. F. We don’t know.” You would have to pick “F” to get a good score, because that’s the current consensus… unless your professor had recently written a paper espousing one of the other answers (see below). “G. None of the above” is the better answer according to the large reptile tree where fenestrasaurs are more closely related to pterosaurs. But each one of the above (A-E) has been proposed by recent authors, not caring if they made sense or not.

Imagine the plight of the poor student in Paleontology 101 today
when he or she asks the professor about that website, “repitleevoluton.com” The professor is going to have to say, “If you want a good grade, you’ll ignore that website and provide the same answers that are in your textbook.” That’s what Dr. Darren Naish reported online. Don’t consider, test or discuss other possibilities. Best to ignore them — if you want to advance in paleontology and get your Masters or PhD.

Take, as an example,
David Hone’s dissertation that was later published in two papers in which he proposed comparing two competing pterosaur origin hypotheses, one by Peters 2000 (Cosesaurus, Sharovipteryx, Longiasquama) and one by Bennett 1996 (Scleromochlus) using the supertree method of analysis (combining several published analyses without actually examining any fossil specimens). Aware that his professor, Michael Benton, had earlier written a paper (Benton 1999) celebrating Scleromochlus as the sister to pterosaurs, Hone decided to delete and diminish the taxa proposed by Peters. He somehow created several typos in the Peters data and then deleted the entire Peters dataset because of those typos (references and the full story here). Then Hone and Benton (2008) gave credit for both competing hypotheses to Bennett while deleting all reference to Peters 2000. As a result, Hone received his PhD, two associated papers (Hone and Benton 2007, 2008) were published and Hone gained the ability to referee pterosaur manuscripts (like mine) submitted to academic journals. I wrote to Dr. Benton about the inconsistencies and leaps of logic between the two parts of their two part paper. His reply was a sheepish, “whoops. : )”

See how it works?
That’s how you crush an opposing hypothesis. And that’s just the tip of the iceberg of current readily solvable problems, as Pterosaur Heresies readers are well aware. No PhD wants to admit he/she was wrong. On some problems consensus will likely never be achieved — because in order to do so all invalid candidate hypothesis writers would have to admit they were wrong.

And that’s just not going to happen.Not without a fight or a dismissal. Let me know if you know of any instances of someone admitting they were wrong (I know of one semi-wrong situation regarding Dr. Padian and his fight with pterosaur tracks). In the origin of snakes, pterosaurs, turtles and dinosaurs there are lots of ‘right’ answers out there, but few challenges to the weaker hypothesis and no one admits to being wrong.

As history tells us, in paleontology it takes decades to turn the boat around. And paleontologists don’t want anyone else, even other paleontologists, solving their mysteries for them… even when solutions are published in the literature.

Thanks for your interest.
I will continue to study and make informed comment on new fossil specimens, (many that haven’t made the textbooks yet). I will throw a spotlight on problems and celebrate solutions as they are verified or not in the large reptile tree. And I encourage you to do the same. If I can do it, anyone can do it.

There are too many paleontologists who
follow matrices, textbooks and papers blindlyand not enough paleontologists who have the balls to say, “Hey, there’s something wrong here.”

We’ll help fix the world of paleontology someday.
Unfortunately, it’s not going to happen this year. After four years of working with the large reptile tree, and improving it, and enlarging it year after year, it still has not been accepted for publication or gained intrigue among basal reptile workers. They don’t like it. It rocks the boat.

ReferencesBennett SC 1996. The phylogenetic position of the Pterosauria within the Archosauromorpha. Zoolological Journal of the Linnean Society 118: 261–308.Benton MJ 1999.Scleromochlus taylori and the origin of the pterosaurs. Philosophical Transactions of the Royal Society London, Series B 354 1423-1446. Online pdfHone DWE and Benton MJ 2007. An evaluation of the phylogenetic relationships of the pterosaurs to the archosauromorph reptiles. Journal of Systematic Palaeontology 5:465–469.Hone DWE and Benton MJ 2008. Contrasting supertree and total evidence methods: the origin of the pterosaurs. Zitteliana B28:35–60.Peters D 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.

Sorry to have to report this, but…
Witton (2015) decided that certain published literature (data and hypotheses listed below) germane to his plantigrade, quadrupedal, basal pterosaur conclusions, should be omitted from consideration and omitted from his references.

Everyone knows, iI’s always good practice
to consider all the pertinent literature. And if a particular observation or hypothesis runs counter to your argument, as it does in this case, your job is to man up and chop it down with facts and data. That could have been done, but wasn’t. Instead, Witton put on his blinders and pretended competing literature did not exist. Unfortunately that’s a solution that is condoned by several pterosaur workers of Witon’s generation.

Not the first time inconvenient data
has been omitted from a pterosaur paper. Hone and Benton (2007, 2008) did the same for their look into pterosaur origins after their own typos cleared their way to delete from their second paper one of the two competing candidate hypotheses.

Witton (2013) and Unwin (2005) did the much the same by omitting published papers from their reference lists that they didn’t like.

Publication
is a great time to show colleagues that you have repeated all competing observations and experiments and either you support or refute them. To pretend competing theories don’t exist just increases controversy and reduces respect.

So, what’s this new Witton paper all about?

From the Witton abstract:“Pterodactyloid pterosaurs are widely interpreted as terrestrially competent, erect-limbed quadrupeds, but the terrestrial capabilities of non-pterodactyloids are largely thought to have been poor (false). This is commonly justified by the absence of a non-pterodactyloid footprint record (false according to Peters 2011), suggestions that the expansive uropatagia common to early pterosaurs would restrict hindlimb motion in walking or running (false), and the presence of sprawling forelimbs in some species (not pertinent if bipedal).

“Here, these arguments are re-visited and mostly found problematic. Restriction of limb mobility is not a problem faced by extant animals with extensive fight membranes, including species which routinely utilize terrestrial locomotion. The absence of non-pterodactyloid footprints is not necessarily tied to functional or biomechanical constraints. As with other fully terrestrial clades with poor ichnological records, biases in behaviour, preservation, sampling and interpretation likely contribute to the deficit of early pterosaur ichnites. Suggestions that non-pterodactyloids have slender, mechanically weak limbs are demonstrably countered by the proportionally long and robust limbs of many Triassic and Jurassic species. Novel assessments of pterosaur forelimb anatomies conflict with notions that all non-pterodactyloids were obligated to sprawling forelimb postures. Sprawling forelimbs seem appropriate for species with ventrally-restricted glenoid articulations (seemingly occurring in rhamphorhynchines and campylognathoidids). However, some early pterosaurs, such as Dimorphodon macronyx and wukongopterids, have glenoid arthrologies which are not ventrally restricted, and their distal humeri resemble those of pterodactyloids. It seems fully erect forelimb stances were possible in these pterosaurs, and may be probable given proposed correlation between pterodactyloid-like distal humeral morphology and forces incurred through erect forelimb postures. Further indications of terrestrial habits include antungual sesamoids, which occur in the manus and pes anatomy of many early pterosaur species, and only occur elsewhere in terrestrial reptiles, possibly developing through frequent interactions of large claws with firm substrates. It is argued that characteristics possibly associated with terrestrially are deeply nested within Pterosauria and not restricted to Pterodactyloidea as previously thought, and that pterodactyloid-like levels of terrestrial competency may have been possible in at least some early pterosaurs.”

Witton writes:“Given that likely pterosaur outgroups such as dinosauromorphs and Scleromochlus bore strong, erect limbs (e.g.,Sereno, 1991; Benton, 1999), it is possible that these early pterosaurs retained characteristics of efficient terrestriality from immediate pterosaur ancestors.”

Wrong as this ‘given’ supposition is, both of the above taxa (dinos and scleros) are bipedal, yet Witton refuses to consider this configuration in basal pterosaurs (for which he claims have no ichnite record).

Figure 1. Witton’s errors with a quadrupedal Preondactylus. For a study on terrestrially, there is little effort devoted to the feet of pterosaurs here. Click to enlarge.

Digitigrady vs. plantigrady
Pterosaur feet come in many shape and sizes. Some have appressed metatarsals. Others spread the metacarpals. These differences were omitted by Witton. Some have a very long pedal digit 5. Others have a short digit 5. These differences were also omitted. Some pterosaurs were quadrupeds (but not like Witton imagines them), others were bipeds (Figs. 1-6). Basal pterosaurs had a butt-joint metatarsi-phalangeal joint, but that just elevates the proximal phalanges, as confirmed in matching ichnites.

Figure 2. Witton’s quadrupedal Dimorphodon. Click to enlarge.

The quadrupedal hypothesis is a good one,
but it really only works in short-clawed plantigrade clades that made quadrupedal tracks on a horizontal substrate. Otherwise a quadrupedal configuration works only on vertical surfaces, like tree trunks, where the trenchant manual claws can dig into the bark. This was omitted by Witton.

Figure 3. Dimorphdon toes and fingers. Here, in color, I added the keratinous sheath over the claws that show how ridiculous it would be for Dimorphodon to grind these into the ground. Better to use those on a vertical tree trunk (figure 2). Click to enlarge.

Pterosaurs were built for speed
whether on the ground or in the air. They were never ‘awkward.’ Remember basal forms have appressed metatarsals, they have more than five sacrals, their ichnites are digitigrade, the tibia is longer than the femur, the bones are hollow, when bipedal the feet plant below the center of balance at the wing root, and some pterodactyloid tracks are bipedal.

Figure 6. Quadrupedal Campylognathoides by Witton (center) with errors noted and compared to bipedal alternatives. The lack of accuracy in Witton’s work borders on cartoonish.

Accuracy trumped by imagination
By the present evidence, Witton has not put in the effort to create accurate and precise pterosaur reconstructions. Rather his work borders on the cartoonish and I suspect the reconstructions have been free-handed with missing or enigmatic parts replaced with parts from other pterosaurs. That should be unacceptable, but currently such shortcuts are considered acceptable by Witton’s generation of pterosaur workers.

The Sordes uropatagium false paradigm gets a free pass
and no critical assessment from Witton. (So far this uropatagium has been observed only in one specimen, Sordes (in which a single uropatagium Witton believes was stretched between the two hind limbs), was shown to be an illusion caused by bone and membrane dislocation during taphonomy. All other pterosaurs and their predecessors have twin uropatagia that do not encumber the hind limbs. The dark-wing Rhamphorhynchus (Fig. 5) is an example of a basal pterosaur with twin uropatagia.

So is Margot Garritsen, a Dutch engineer and Stanford professor who leads a team intent on building a flying pterosaur based on Paul Sereno’s ornithocheirid from the Sahara. They were counting on greater success with lighter materials and a more accurate wing movement for flight control.

Dino Frey (Natural History Museum of Karlsruhe) is featured with a giant ‘wing bone’ from Israel having only a cylindrical body without articular ends. Looks to be about 8 inches in diameter, more than 8 feet long (60-foot, 18 m wingspan or twice the size of Quetzalcoatlus). It made the news here and here. Giant pterosaurian footprints from Mexico appear to confirm the size, all discovered prior to 2005, still not published.

On that note:Mark Witton reported on the DML in 2008, “However, subsequent reappraisals of the alleged discoveries suggested that the footprints belong to a large theropod dinosaur and the ‘wing bone’ is, in fact, a particularly large piece of fossil wood (E. Frey, pers. comm. 2007), suggesting claims of 20 m flying reptiles were somewhat premature.”

Yes, even PhDs sometimes make mistakes. And later in the video the giant pterosaur ‘bone’ is confirmed as wood. Other problems you’ll no doubt recognize. Lot’s of bad and speculative propaganda here.

Some good data from Kevin Padian on pterosaur landings. You can see an earlier animation here, but the video has a new one in 3-D.

“Palaeontology [online] is a website covering all aspects of palaeontology. The site is updated with articles about the cutting edge of research, by the researchers themselves. These are usually written by experts in the field, but are aimed at non-specialists. Articles vary widely in their content: some serve as an introduction to palaeontological or interdisciplinary fields, while others outline events in the history of palaeontology. Some contributions include summaries of recent findings and advances in rapidly evolving disciplines, and some focus on a particular geographic region or time period. Finally, some of our articles are based on the experience of being a palaeontologist – what life and work is really like as a fossil worker. Our online format allows researchers to explain their work with the aid of an unlimited number of figures and videos.”

Commissioning editors (who are responsible for inviting contributions and overseeing the website) are:

The pterosaur page was written by Dr. David Hone, who states, “The origins and the relationships of the pterosaurs have long been contentious, although a consensus is forming on both issues. Often confused with dinosaurs, pterosaurs are members of their own clade, but are close relatives of their more famous cousins.

Over the years, palaeontologists have hypothesized that pterosaurs originated from various parts of the reptile evolutionary tree. Very early researchers considered them to be the ancestors of birds or even bats, and for a long time it seemed that they were probably basal archosaurs (the clade that contains dinosaurs, birds, crocodilians and some other groups). More recently evidence has begun to stack up that they are a separate group to the dinosauromorphs (dinosaurs and their closest relatives) but that the two groups evolved from a common ancestor. Most researchers now support this position. This makes pterosaurs reasonably close relatives to birds, but they are not bird ancestors as is sometimes wrongly reported.”

Well, par for the course…
Sad to see when there actually is a verifiable better relationship out there, but then that would involve actually acknowledging the literature (Peters 2000, 2002, 2007, 2009, 2011) and/or testing candidates one vs. another. But nobody wants to do this without fudging the data or reducing the inclusion set. It’s time to either recognize the literature or argue with it. The large reptile tree found a long line of pterosaur ancestors between Ichthyostega and Longisquama. Almost any one will do, as we learned earlier with turtles and pterosaurs.